Invasive rodents have multiple indirect effects on seabird island invertebrate food web structure.

Burrowing seabirds that nest on islands transfer nutrients from the sea, disturb the soil through burrowing, damage tree foliage when landing, and thereby modify the surface litter. However, seabirds are in decline worldwide, as are their community- and ecosystem-level impacts, primarily due to invasive predatory mammals. The direct and indirect effects of seabird decline on communities and ecosystems are inherently complex. Here we employed network analysis, as a means of simplifying ecological complexity, to better understand the effects seabird loss may have on island invertebrate communities. Using data on leaf litter communities, we constructed invertebrate food webs for each of 18 offshore oceanic islands in northeastern New Zealand, nine of which have high seabird densities and nine of were invaded by rats. Ten network topological metrics (including entropy, generality, and vulnerability) were compared between rat-invaded and uninvaded (seabird-dominant) islands. We found that, on rat-invaded islands, the invertebrate food webs were smaller and less complex than on their seabird-dominated counterparts, which may be due to the suppression of seabird-derived nutrients and consequent effects on trophic cascades. This decreased complexity of food webs due to the presence of rats is indicative of lower ecosystem resistance via lower trophic redundancy. Our results show that rat effects on island ecosystems are manifested throughout entire food webs, and demonstrate how network analysis may be useful to assess ecosystem recovery status as these invaded islands are restored.

Invasive mammal eradication on islands results in substantial conservation gains.

More than US$21 billion is spent annually on biodiversity conservation. Despite their importance for preventing or slowing extinctions and preserving biodiversity, conservation interventions are rarely assessed systematically for their global impact. Islands house a disproportionately higher amount of biodiversity compared with mainlands, much of which is highly threatened with extinction. Indeed, island species make up nearly two-thirds of recent extinctions. Islands therefore are critical targets of conservation. We used an extensive literature and database review paired with expert interviews to estimate the global benefits of an increasingly used conservation action to stem biodiversity loss: eradication of invasive mammals on islands. We found 236 native terrestrial insular faunal species (596 populations) that benefitted through positive demographic and/or distributional responses from 251 eradications of invasive mammals on 181 islands. Seven native species (eight populations) were negatively impacted by invasive mammal eradication. Four threatened species had their International Union for the Conservation of Nature (IUCN) Red List extinction-risk categories reduced as a direct result of invasive mammal eradication, and no species moved to a higher extinction-risk category. We predict that 107 highly threatened birds, mammals, and reptiles on the IUCN Red List-6% of all these highly threatened species-likely have benefitted from invasive mammal eradications on islands. Because monitoring of eradication outcomes is sporadic and limited, the impacts of global eradications are likely greater than we report here. Our results highlight the importance of invasive mammal eradication on islands for protecting the world's most imperiled fauna.

Burrowing seabird effects on invertebrate communities in soil and litter are dominated by ecosystem engineering rather than nutrient addition.

Vertebrate consumers can be important drivers of the structure and functioning of ecosystems, including the soil and litter invertebrate communities that drive many ecosystem processes. Burrowing seabirds, as prevalent vertebrate consumers, have the potential to impact consumptive effects via adding marine nutrients to soil (i.e. resource subsidies) and non-consumptive effects via soil disturbance associated with excavating burrows (i.e. ecosystem engineering). However, the exact mechanisms by which they influence invertebrates are poorly understood. We examined how soil chemistry and plant and invertebrate communities changed across a gradient of seabird burrow density on two islands in northern New Zealand. Increasing seabird burrow density was associated with increased soil nutrient availability and changes in plant community structure and the abundance of nearly all the measured invertebrate groups. Increasing seabird densities had a negative effect on invertebrates that were strongly influenced by soil-surface litter, a positive effect on fungal-feeding invertebrates, and variable effects on invertebrate groups with diverse feeding strategies. Gastropoda and Araneae species richness and composition were also influenced by seabird activity. Generalized multilevel path analysis revealed that invertebrate responses were strongly driven by seabird engineering effects, via increased soil disturbance, reduced soil-surface litter, and changes in trophic interactions. Almost no significant effects of resource subsidies were detected. Our results show that seabirds, and in particular their non-consumptive effects, were significant drivers of invertebrate food web structure. Reductions in seabird populations, due to predation and human activity, may therefore have far-reaching consequences for the functioning of these ecosystems.

Surviving in a semi-marine habitat: Dietary salt exposure and salt secretion of a New Zealand intertidal skink.

Species that inhabit marine and intertidal ecosystems face osmoregulatory challenges, risking dehydration and increased ion concentrations in the body. Lizards need to either tolerate or regulate these increased ion concentrations. In this study, we aim to understand how Suter's skink (Oligosoma suteri), an intertidal skink restricted to shoreline habitats, is able to cope with the physiological stress of living in an extreme salt environment. We determined the diet, prey species' salt content, and nasal and cloacal salt excretion on Rangitoto and Motutapu Islands in northern New Zealand, where the skinks have contrasting access to terrestrial invertebrates. Analysis of stable isotopes suggests inter- and intra-population variability in Suter's skink diets. Intertidal invertebrates under washed up seaweed appear to compose a major part of the diet of the Rangitoto population, while the Motutapu population showed evidence for a mixed diet of terrestrial and intertidal invertebrates. Sodium content of prey species decreased with an increasing distance from the seawater. Field secretion of cations through nasal glands consisted primarily of Na(+), which is consistent with other marine and intertidal species. Sodium was also the primary cation in urine. In contrast, fecal cations consisted primarily of K(+). This first study into the salt secretion of an intertidal skink species provides evidence of Suter's skink's plasticity in secreting excess ions through nasal salt glands. This likely enables it to deal with the challenges of living in a semi-marine habitat.

Moving house: long-term dynamics of corticosterone secretion are unaltered in translocated populations of a rare reptile (the tuatara, Sphenodon punctatus).

Translocations are an important conservation tool used to restore at-risk species to their historical range. Unavoidable procedures during translocations, such as habitat disturbance, capture, handling, processing, captivity, transport and release to a novel environment, have the potential to be stressful for most species. In this study, we examined acute and chronic stress (through the measurement of the glucocorticoid corticosterone) in a rare reptile (the tuatara, Sphenodon punctatus). We found that: (i) the acute corticosterone response remains elevated during the initial translocation process but is not amplified by cumulative stressors; and (ii) the long-term dynamics of corticosterone secretion are similar in translocated and source populations. Taken together, our results show that translocated tuatara are generally resistant to cumulative acute stressors and show no hormonal sign of chronic stress. Translocation efforts in tuatara afford the potential to reduce extinction risk and restore natural ecosystems.

Modulation of corticosterone secretion in tuatara (Sphenodon punctatus): Evidence of a dampened stress response in gravid females.

Baseline and stress response glucocorticoid (GC) secretion can be modulated by individuals to support activities and physiological functions connected with reproduction (migration, mating, oviposition and/or parturition, care of young). Corticosterone (CORT) is the primary GC in reptiles and, in accordance with other vertebrates, an adrenocortical stress response is observed. Modulation of CORT secretion occurs in several reptile species, such that elevated baseline CORT concentration and/or a dampened CORT response are common during reproductive life-history events. We investigated CORT secretion after 24 h capture-restraint in the oviparous tuatara (Sphenodon punctatus), the last living rhynchocephalian, and tested whether gravid females have a dampened CORT response compared with non-gravid females. We also included males as a comparison. We confirmed that gravid females have significantly higher baseline plasma CORT concentrations than non-gravid females, suggesting increased CORT secretion during nesting. Furthermore, we found that gravid females exhibit a dampened CORT response compared to non-gravid females and males. Our results demonstrate that female reproductive condition influences CORT secretion in tuatara, and suggest that gravid females modulate CORT secretion during nesting to maintain homeostasis, effectively increasing chances of reproductive success and promoting overall fitness.

Drivers of seabird population recovery on New Zealand islands after predator eradication.

Eradication of introduced mammalian predators from islands has become increasingly common, with over 800 successful projects around the world. Historically, introduced predators extirpated or reduced the size of many seabird populations, changing the dynamics of entire island ecosystems. Although the primary outcome of many eradication projects is the restoration of affected seabird populations, natural population responses are rarely documented and mechanisms are poorly understood. We used a generic model of seabird colony growth to identify key predictor variables relevant to recovery or recolonization. We used generalized linear mixed models to test the importance of these variables in driving seabird population responses after predator eradication on islands around New Zealand. The most influential variable affecting recolonization of seabirds around New Zealand was the distance to a source population, with few cases of recolonization without a source population ≤25 km away. Colony growth was most affected by metapopulation status; there was little colony growth in species with a declining status. These characteristics may facilitate the prioritization of newly predator-free islands for active management. Although we found some evidence documenting natural recovery, generally this topic was understudied. Our results suggest that in order to guide management strategies, more effort should be allocated to monitoring wildlife response after eradication.

Responses of Tuatara (Sphenodon punctatus) to removal of introduced Pacific rats from islands.

Invasive mammalian predators such as rats are now widespread on islands, but hypotheses about their effects have rarely been tested. Circumstantial evidence from New Zealand indicates that, when introduced to islands, Pacific rats (Rattus exulans) have negative effects on endemic plants, invertebrates, birds, and reptiles, including the tuatara (Sphenodon punctatus). We tested the effects of Pacific rats on tuatara by comparing the demographic structure and body condition of tuatara populations on three islands before and after removal of rats and on a fourth island where rats remained. In the presence of rats, juvenile tuatara constituted on average 0-5% of the sample tuatara populations. When Pacific rats were removed after at least 200 years' occupancy, the proportion of juvenile tuatara increased 3.5- to 17-fold and body condition of adult males and females also improved (sometimes dramatically). We predict that, unless Pacific rats are removed from Taranga Island, the tuatara population will collapse because of low population density and the lack of juvenile recruitment. Our results demonstrate that when invasive species exert subtle effects on recruitment and body condition, the effects on populations of long-lived endemic species may only become apparent long after the invasion.

Above- and below-ground impacts of introduced predators in seabird-dominated island ecosystems.

Predators often exert multi-trophic cascading effects in terrestrial ecosystems. However, how such predation may indirectly impact interactions between above- and below-ground biota is poorly understood, despite the functional importance of these interactions. Comparison of rat-free and rat-invaded offshore islands in New Zealand revealed that predation of seabirds by introduced rats reduced forest soil fertility by disrupting sea-to-land nutrient transport by seabirds, and that fertility reduction in turn led to wide-ranging cascading effects on belowground organisms and the ecosystem processes they drive. Our data further suggest that some effects on the belowground food web were attributable to changes in aboveground plant nutrients and biomass, which were themselves related to reduced soil disturbance and fertility on invaded islands. These results demonstrate that, by disrupting across-ecosystem nutrient subsidies, predators can indirectly induce strong shifts in both above- and below-ground biota via multiple pathways, and in doing so, act as major ecosystem drivers.

Intercepting the first rat ashore.

A single Norway rat released on to a rat-free island was not caught for more than four months, despite intensive efforts to trap it. The rat first explored the 9.5-hectare island and then swam 400 metres across open water to another rat-free island, evading capture for 18 weeks until an aggressive combination of detection and trapping methods were deployed simultaneously. The exceptional difficulty of this capture indicates that methods normally used to eradicate rats in dense populations are unlikely to be effective on small numbers, a finding that could have global implications for conservation on protected islands.